What big teeth you have…

One of Charles R. Knight’s paintings of Smilodon fatalis, this one menacing a giant sloth stuck in tar (off panel).

There are few fossil mammals that are as impressive the saber-toothed cat Smilodon fatalis, but despite it’s fearsome dentition some recent new reports have suggested it was more of a pussycat when it came to bite strength. This seems to be counter-intuitive; how could such an impressive animal be associated with the term “weak”? Part of it has to do with word choice, but the larger issue has to do with the fact that the bite of Smilodon wasn’t as strong as that of some other carnivores (extinct and extant), as well as dentition and feeding ecology. This issue goes far beyond just one genus or species, however, as Smilodon was only one of many genera that bore massive canines. In fact, huge “sabers” have evolved over-and-over again in the mammalian lineage (see this post and also this post for information about the cat-like ones), including the famous fangs of the machairodontine felids (saber-toothed cats) and their look-alike nimravid relatives.

Lateral, anterior, and dorsal views of the herbivore Uintatherium (Note the prominent canines). From Marsh, O.C. “The Fossil Mammals of the Order Dinocerata.” The American Naturalist, Vol. 7, No. 3. (Mar., 1873), pp. 146-153

Skull of another member of the Dinocerata; “Loxolophodon cornutus” (today known as Eobasileus cornutus). Again, note the prominent canine. From Cope, E.D. “The Amblypoda (Continued).” The American Naturalist, Vol. 19, No. 1. (Jan., 1885), pp. 40-55.

Although this post will primarily be concerned with the great “sabercats,” large, dagger-like canine teeth having been evolved multiple times by many different unrelated animals during the course of life on earth. In some herbivorous creatures, like the extinct Uintatherium and even in the extant Musk Deer, the fangs reflect sexual dimorphism and probably sexual selection, but the sharp teeth don’t seem to have a prominent function in mastication or processing of food. Likewise, large canine teeth are present in living baboons (Papio sp.), and the sexual dimorphism exhibited between the dental equipment of the males and the smaller canines of the females has long been noted (males often yawn to show off their canines, the size of their teeth being very intimidating indeed). Do the same considerations of sexual selection and dimorphism hold true for the saber-toothed cats, too? Unfortunately, fossil evidence does not always allow comparisons of the two sexes, but extant big cats and some death-trap sites have provided some information to work with. From Salesa, et al. (2006);

Among the Carnivora, sexual dimorphism is more marked in canine size than in other dental features or skull size, and these differences can be related to the breeding system. Species in which a male defends a group of females tend to be more dimorphic than those with monogamous pairs or groups of males and females. Felids are dimorphic animals, but mainly in reference to body size, with the mane of male lions being a unique example of morphological variation between sexes among the family.

This makes sense; if a male keeps a harem of females and has to defend it from other males, the species is more likely to exhibit sexual dimorphism than not. In cats, however, it seems to be more about body size (and possibly characters that wouldn’t fossilize in extinct species) than about tooth size (which would serve important other functions, so any sexual selection would be mitigated by natural selection), although we can’t be sure of this being that there are no living sabercats to study. Personally, I think there could be a sexual-selection component in some groups, but the saber-canine is so prominent in so many extinct felids and nimravids that it is extremely doubtful that all the lineages converged on similar tooth structures because of sexual selection/dimorphism, the functional advantage of larger teeth likely coming first. A lack of sexual dimorphism when considering morphology as a whole, however, may suggest a more solitary lifestyle where territories may or may not overlap are maintained and direct competition for females is not as fierce, especially since the females move through territories rather than living with a male. Such a strategy may have been employed by the late Micoene sabercat Paramachairodus ogygia. Salesa, et al. (2006), working with an assemblage made up of many of the more basal felids, have even been able to come up with a hypothesis about life history of the ancient animals based upon their finds in Spain;

[T]he probable territorial behaviour for Par. ogygia would be very similar to that of jaguars, in which males defend large, overlapping territories that include smaller territories of several females. This model is similar to that of the leopard, but in this species male territories never overlap, which could explain the different sexual dimorphism index of this species with respect to Par. ogygia and jaguar…

So, if Par. ogygia behaved more like jaguars and leopards than lions, the presence of juveniles in the trap would be highly improbable, as is the case. But in addition to the scarcity of juveniles, the sample from Batallones-1 has another interesting feature: it is mostly composed of young adults, that is, individuals with the complete permanent dentition, but without any trace of wear. These animals, which would have recently become independent of their mothers, would not as yet have had any territory, moving instead through the ranges of other adults and being more easily attracted by an easy meal, such as carrion. This age distribution therefore suggests that the sample of Par. ogygia trapped in Batallones-1 corresponds to that fraction of non-resident young individuals, both males and females, which were in a phase of dispersion. In the case of leopards, such individuals are more daring – or less cautious – than adults, and they have been seen crossing rivers in spate, whereas resident adults only cross at times of lower water. It has also been noticed that among these individuals, males are even more inclined to make these incursions than females, which remain longer with the mother, especially if there is good availability of food. If this pattern of dispersion behaviour applied to the young adults of Par. ogygia, it is likely that they were trapped in Batallones-1 more often than the resident adults.

Saber-Toothed Felid and Nimravid diversity (click for a larger image). From Emerson, S.B., and Radinsky, L. “Functional Analysis of Sabertooth Cranial Morphology.” Paleobiology, Vol. 6, No. 3. (Summer, 1980), pp. 295-312.

While the life histories of extinct mammalian carnivores are interesting in and of themselves, it is the teeth and terrifying bite of the sabercats that we are most concerned with here. Smilodon is the celebrity of saber-toothed cats, but the fossil record preserves a wide diversity of carnivores with large canine teeth, and even within the larger groupings there are even more subdivisions, the skulls of saber-toothed felids being widely variable. As discussed in the background material, nimravids are saber-tooth look-alikes that diverged from a common ancestral line earlier than the carnivores that would give rise to Smilodon, but the two lines are still closely related and have undergone parallel evolution. There is still some reshuffling of taxa going on and the true evolutionary history/affinities of many of the forms is still being worked out, but most forms you’re likely to see grouped together at a museum fall into either the nimravid or felid camps. The focus of this essay, however, will be on felids, and although they are often discussed along with their nimravid cousins the larger amount of work has been done on the felids and so we must leave the nimravids.

With the felids, then, there seem to be three kinds of sabercat that hint at differing predatory tactics, prey, and habitat. Indeed, evolution did not create carbon copies of the same creature, barring life from becoming adapted to varying circumstances; there is more variety than would be first assumed if we based all our research on the presence of prominent canines. Instead, there seem to be three “ways of being” a saber-toothed cat, as outlined by Martin, et al.;

Saber-toothed carnivores… have been divided into two groups: scimitar-toothed cats with shorter, coarsely serrated canines coupled with long legs for fast running, and dirk-toothed cats with more elongate, finely serrated canines coupled to short legs built for power rather than speed. In the Pleistocene of North America, as in Europe, the scimitar-cat was Homotherium; the North American dirk-tooth was Smilodon. We now describe a new sabercat from the Early Pleistocene of Florida [Xenosmilus], combining the scimitar-tooth canine with the short, massive limbs of a dirk-tooth predator. This presents a third way to construct a saber-toothed carnivore.

Xenosmilus hodsonae, Homotherium cf. crenatidens, and Homotherium serum. From Martin, L.D., Babiarz, J.P., Naples, V.L., and Hearst, J. “Three Ways To Be a Saber-Toothed Cat.” Naturwissenschaften, Vol. 87, No. 1 (Jan. 2000), pp. 41-44

As Martin notes, there appears to be a number of adaptational “trade offs” that sabercats in North America and Europe were subject to; fast-moving gracile forms had shorter sabers, but stouter and more powerful forms had the longer, more laterally flattened canine teeth. The “third way” that combined characters from both groups was exemplified by Xenosmilus (which Martin, et al. say would have seemed more like a bear than a cat, despite actual evolutionary relationships to the contrary). Still, leaving the overall structure of the body aside for a moment, the arrangement and sizing of the teeth of the different groups can be very telling. Martin, et al. again lay out what the usefulness of the differing tooth arrangements;

When biting, the long sabers of dirk-toothed cats may have cut parallel slits for some distance before the relatively smaller incisors could be applied. In scimitar-toothed cats the shorter canines and longer incisors worked more as a unit, first cutting parallel slits with the canines, immediately followed by the incisor arc removing the strip of flesh. Such a large open wound would have bled profusely, traumatizing the victim. If the incisors and canines acted in unison, the torsional forces on individual teeth would have been reduced, resulting in fewer restrictions on bite placement. In felids the size of the sagittal crest is directly proportional to the forces exerted by the temporalis musculature. Scimitar-toothed cats have a sagittal crest that is generally less pronounced than that in their dirk-toothed contemporaries. In a modification of the typical scimitar-tooth condition, the new cat from Florida exhibits both an elongated sagittal crest and an enlarged temporalis muscle that would have permitted a stronger bite.

While such a passage might not seem significant at first, it shows that there is more going on in a sabercat’s skull that is important to biting than just the size or shape of the canines. The placement of the incisors, for instance, seem to make a difference in biting strategy and force, dirk-toothed cats like Smilodon exhibiting a condition where the incisors are out forward of the canines. When this is taken into account, as well as the length of the canines, it seems that the canines would slash for quite some distance before the incisors could be used at all in comparison to the scimitar-toothed sabercats, the placement of the incisors in scimitar-tooths seemingly strengthening the biting teeth at the front of the jaw. The sagittal crests of these creatures should also be taken into account, such structures giving students of paleontology an indication of how carnivores (or herbivores, in the case of gorillas) have been adapted to achieve higher bite forces. Such ridges atop the skull for muscle attachment are not unique to sabercats, however, and there are some animals that have taken the structure to even greater extremes;

The extinct “bear dog” Amphicyon at the AMNH. Note the size of the sagittal crest, the reduction of the bony enclosure around the eyes, and the large holes on the side of the skull for jaw muscle attachment.

The extinct “saber-toothed” creodont Hyaenodon at the AMNH. Again, note the sagittal crest, reduction of bone enclosure around the eye, and the large canines.

The skull of the nimravid Hoplophoneus on display at the AMNH. Note the size of the canines and sagittal crest in comparison with Hyaenodon and Amphicyon.

The skull of Smilodon on display at the AMNH.

The skull of the marsupial predator Thylacoleo at the AMNH. Note the large openings on either side of the skull for the jaw muscles.

Ventral view of the skull of Thylacoleo. From E.D. Cope’s “The Tertiary Marsupialia” in The American Naturalist, Vol. 18, No. 7. (Jul., 1884), pp. 686-697.

Looking at the various groups, all show adaptations that increase the amount of available muscle attachment to achieve more powerful bites, modifying the skull in two ways. First, a sagittal crest (as already discussed) is often present to some degree, often being greater in omnivores or bone-crushing carnivores as they require greater forces to crack hard foods (although recent research by Wroe, et al. suggest that bone crushers like Spotted Hyena might not have the highest bite forces). Likewise, the holes between the skull and cheek bones are often enlarged or widened (the extreme of this group being Thylacoleo), the more muscle that can pass from lower jaw to skull being directly correlated to bite strength. What is interesting about sabercats, when considering these factors, is that they seem to be in the middle. They don’t exhibit adaptations of the skull to the extreme as in Amphicyon or Thylacoleo, but they still exhibit changes allowing for powerful bites (strong enough to kill and consume prey, at least). The trend is obvious and has not been missed by reseachers, and Emerson says the following about it;

With enlargement of upper canines, skulls of paleofelid, neofelid, marsupial and, as far as the record shows, creodont sabertooths were remodeled in similar ways. This evolutionary convergence in cranial morphology is not surprising, since most of the modifications relate to allowing increased gape while retaining bite strength at the carnassial. Those are factors essential for all sabertooths, and the possible ways to achieve them, starting from a generalized mammalian cranial morphology, are limited…

Why did sabertooth specializations evolve so many times? Their multiple evolution, plus the fact that several species of sabertoothed felids existed for most of the history of the family (from about 35 Myr to about 15,000 yr BP) suggest that sabertooth canines provided an effective alternative to the modern carnivore mode of killing prey

The skull of the saber-toothed cat Megantereon. Like in Smilodon, not how the incisors jut out (as well as the overly large nasal opening in this genus).

The basic mechanics of the skull just discussed gives researchers clues as to how sabercats could have killed their prey, but reconstructing ancient predator/prey interactions with no exact modern equivalent is difficult. Indeed, debate has gone on for years as to how sabercats used their teeth to bring down prey (see Simpson’s paper), either by stabbing, cutting, slicing, or even (as silly as it may seem) by crushing. What does seem apparent today, however, is that the canines of the sabercats were relatively delicate, and it would be unwise to fully sink them into a struggling animal as they may easily be broken off. Even if such an attempt to deeply puncture a prey item was not undertaken, biting full-force into bone could have also easily damaged teeth (or even broken them off), making it unlikely that sabercats jumped onto the back of their prey and tried to sink their teeth into the back of the prey’s skull like some modern cats. Recent research has even shown that the skull of Smilodon was ill-suited to handle stresses associated with struggling prey when compared to the skull of a lion, and I wonder how often individual Smilodon perished because of stresses associated with taking down prey if the victim was not brought down and killed quickly. Indeed, it seems that the long teeth were better suited to slicing soft flesh, i.e. cutting open the belly of prey or slicing open the throat, rather than piercing rough hides and ramming through bone.

Skulls (mandibles not pictured) of 4 “saber-toothed” mammals from “The Function of Saber-Like Canines in Carnivorous Mammals” by G.G. Simpson, American Museum Novitiates, August 4, 1941. Pictured are A) Machairodus (felid), B) Hoplophoneus (nimravid), C) Smilodon (felid), and D) Thylacosmilus (marsupial).

As just discussed in terms of tooth and skull stressed, many factors of life history, behavior, and morphology of extant big cats and sabercats might be similar, but the massive canines of the extinct group seem to infer a different killing strategy, and there is no reason to assume that they were like modern big cats in every respect. Salesa, et al. sums it up this way;

Extant felids kill small animals by biting on the nape or directly on the skull, using their rounded-section canines, but if any sabre-toothed cat tried to do this they would have risked breaking the laterally flattened upper canines. For this reason, it is more probable that they developed some behavioural mechanism to minimize that risk, such as ignoring prey below a given size. It is likely that machairodontines developed this ethological trait early in their evolution, and so narrowed their prey size range in comparison with that of felines, which hunt both large and small animals. This high specialization has been pointed out as one of the possible reasons for the gradual decline and final extinction of the sabre-toothed cats in the Pleistocene… The development of this strategy was probably the key reason for the sabre-tooted cats becoming the dominant predators in the land mammal faunas from the Late Miocene to Late Pleistocene.

It might not immediately make sense that felids with fragile teeth would specialize in eating large prey, but that is whale the fossil evidence (as we currently understand it) infers. While the smallest prey would pose no problems (outside of not being a fully satisfying meal), but medium sized prey with smaller areas of soft flesh (like the stomach and neck) would potentially be more dangerous and a more exact bite would be needed to prevent damage to the teeth and skull. Hence, it seems that the slashing and ripping of soft tissue in larger animals was the main method of killing prey (after it had been brought down or slowed by blood loss), taking hypercarnivory to an even more specialized extent.

An Amur Leopard yawns. Note the relatively small (but still fearsome) canines of the upper and lower jaw.

What, then, of a smaller living cat, the Clouded Leopard (Neofelis nebulosa and N. diardii), which has been heralded as a modern analog of sabercats? As Christiansen notes, Clouded Leopards are a bit bizarre, and it is incorrect to call them “small” big cats or modern sabercats, the genus showing a number of convergences with extinct forms while remaining distinct from the famed genus Panthera;

The skull morphology of the clouded leopard sets it apart from other extant felids, and in a number of respects it approaches the morphology of primitive sabertooths. This indicates convergence of several characters in machairodontine felids and the clouded leopard, mainly as adaptations for attaining a large gape. This raises doubts about the characters hitherto considered as distinguishing sabertoothed from nonsabertoothed predators…

Clearly, Neofelis and the sabertooths independently evolved a suite of the same specializations for the same overall purpose of attaining a large gape, a prerequisite for efficient jaw mechanics with large canines, but the reasons for evolving these characters need not have been similar. Based on analyses of lower jaw bending moments and inferred resistance to mechanical loadings, Therrien (2005) suggested that Neofelis could be at the beginning of a new sabertooth radiation. Such claims are difficult to test, however, since the extant sister taxon to Neofelis (Panthera) shares none of its sabertoothed characters, and the fossil record provides no clues of felids closer to Neofelis than Panthera. At present, however, there is little evidence to suggest that Neofelis can be regarded as an “extant sabertooth,” although it clearly shares a number of characters with them that are absent in other extant felids. On the other hand, it cannot be regarded as simply an intermediate between large and small felids, as normally assumed. The presence to some extent of characters normally ascribed to sabertooths in Neofelis raises doubts about their functional and evolutionary significance in primitive machairodonts such as Nimravides or Paramachairodus, hitherto the only reasonably well-known primitive machairodont. Such animals need not have shared the presumed functional skull morphology of later, more derived sabertooths and are perhaps not to be regarded as “sabertoothed” at all, if by sabertoothed is implied animals functionally significantly different from extant felids.

Again, this shows a convergence of functional morphology despite existing evolutionary relationships, many felids being adapted in similar ways. As stated previously, the large canines of saber-toothed predators required the animals to open their jaws wide but also narrowed their predatory niche to some extent. Likewise, various tests seem to show that the bite of sabercats like Smilodon was “weak,” with news reports often relating that the terrible felids were more like big housecats when compared to living big cats. This is a mistake (and it would be a grave one for anyone ever to cross a sabercat), born of a lack of recognition that bite forces exist on a continuum and are related to a number of factors and cannot simply be deemed “weak” or “strong” without further comment. Christiansen relates the bite force of Smilodon as such;

[A]lthough large sabertooths such as Smilodon and Homotherium had weaker bite forces than lions or tigers, their bite forces were broadly comparable to those of jaguars and large leopards, and, thus, cannot be claimed to have been “weak”. Lower bite forces at any given body size were probably evolutionarily possible owing to a marked contribution from the upper cervical musculature to the killing bite, which… was absent in Neofelis and primitive machairodonts such as Paramachairodus. Thus, bite force analysis may constitute a hitherto overlooked parameter in evaluating whether or not primitive machairodonts such as Paramachairodus or Nimravides really did employ a canine shear bite with a marked contribution from the cervical muscles to subdue prey, or killed in a manner similar to extant felids, which requires a stronger killing bite…

In many Plio-Pleistocene communities predator competition was more severe than today, and a sabertooth killing mode could be a way of ensuring faster kill rates, since a throat shear-bite most likely would kill prey faster than a throttling throat bite, common in extant pantherines. In lions, it can take up to 13 minutes to kill large prey, and in such cases the prey is frequently killed by disemboweling by other pride members. In the cheetah a suffocation bite can take even longer to kill prey. Carcass theft and feeding competition is very common among extant large, sympatric predators, and a faster kill mode could be a way of reducing the risk of carcass theft from competing predators. In many large predators with sympatric competitors, rapid consumption can be a way of reducing the risk of carcass theft, and this would most likely have been accentuated in past ecosystems with more intense large predator competition. Accordingly, the morphology and behavior of extant predators need not reflect the circumstances to which they became adapted when they evolved. More intense competition could accelerate the evolution of a sabertooth morphology…

This passage reflects the problems with reconstructing bite forces and predation techniques of extinct creatures; more is involved than just the opening and closing of the jaw. The neck muscles of many sabercats (except in some of the more basal members, as noted) likely contributed to the strength of the bite in a way that’s not directly testable today. Likewise, the killing technique of sabercats might not have required a bite as strong as a modern-day tiger, as in a land filled with other predators, it might simply take too long to try and suffocate a prey animal or bite through the back of their skull. Disemboweling or tearing out the throat of the prey item, by contrast, is a much quicker way to do large amounts of damage but it seems that it would require teamwork, solitary extant big cats often opting for a killing neck bite when the prey has been brought down. Even if this is eventually shown to be incorrect, it should be remembered that bite strength is not everything; despite its large size, the Great White Shark (Carcharadon carcharias) has a relatively weak bite, but it makes up for it with heavily serrated teeth, force of impact when attacking prey, and side-to-side head shaking to saw through its food. Crocodilians, by contrast, have very strong bite forces but they don’t saw through prey or chew, the emphasis being holding on to struggling prey and drowning it before ripping it apart. Such considerations bring us to another point mentioned above in our discussion of scimitar-tooths vs. dirk tooths in that the famous dirk-toothed cats like Smilodon were more powerfully built, seemingly focusing on bringing a large animal down to the ground and then delivering devastating bites once the stomach and neck were exposed (a process that would be made easier by groups working together, as seen in modern examples like lions bringing down giraffes or elephants).

A group of lions brings down a giraffe.

A group of lions brings down an elephant.

A new paper, just out in PNAS, does take the powerful neck muscles of Smilodon into account, however, and the information from the new models appear to corraborate the modern understanding of a felid that captured and killed prey in a way quite different from Panthera. From McHenry, et al.;

Our results demonstrate that bite force driven by jaw muscles was relatively weak in S. fatalis, one-third that of a lion (Panthera leo) of comparable size, and its skull was poorly optimized to resist the extrinsic loadings generated by struggling prey. Its skull is better optimized for bites on restrained prey where the bite is augmented by force from the cervical musculature. We conclude that prey were brought to ground and restrained before a killing bite, driven in large part by powerful cervical musculature. Because large prey is easier to restrain if its head is secured, the killing bite was most likely directed to the neck. We suggest that the more powerful jaw muscles of P. leo may be required for extended, asphyxiating bites and that the relatively low bite forces in S. fatalis might reflect its ability to kill large prey more quickly, avoiding the need for prolonged bites.

Hunting isn’t the only aspect of sabercat predation that seems to have differed from modern carnivores; they way they ate (and what they ate) is somewhat at variance with modern forms, as well. As is apparent at this point, the contact of the canines with bones would have been avoided, and it seems that the hard parts of the skeleton would have been avoided when a sabercat was consuming it. This could differ among different groups (perhaps some of the shorter-toothed forms not being so finicky about bone), but research into microwear patterns on teeth of Smilodon don’t seem to match with wear patterns of any living carnivores, suggesting a different dietary preference. It could be hypothesized, then, that creatures like Smilodon primarily consumed the soft parts of the carcass or what could be removed without too much damage to the teeth, and it should be remembered that living big cats often do not eat every part of the skeleton. Some, like cougars, have favored parts that they eat but end up leaving as much as 40% of the carcass behind. Other predators, especially bone-crushing ones, could take advantage of the leftovers, although the felids might have had to eat quickly as some of their osteophagus competitors may not have been patient (and, in fact, lions and hyenas often fight over kills and steal them from each other today).

Given all the prior considerations, it now seems that sabercats specialized in bringing down relatively large prey down quickly (some likely working in groups to do so), killing the victims by slashing open their stomachs or slicing through the blood vessels of the neck. This would be a much messier, but quicker, method than employed by living big cats, although the limitation of food sources likely caused in the eventual downfall of sabercats. Hypercarnivory can be a dangerous adaptive path to go down, and cats are clearly the most meat-dependant of the Carnivora, but it seems that extinct forms took their dental and dietary specialization above and beyond what is seen today. The price paid for such adaptations ended up being extinction, but given how many times they have shown up in the history of life on this planet, someday there may again be a saber-toothed predator stalking the shadows.

References;

Anyonge, W. “Microwear on Canines and Killing Behavior in Large Carnivores: Saber Function in Smilodon fatalis” Journal of Mammalogy, Vol. 77, No. 4 (Nov., 1996), pp. 1059-1067

Christiansen, P. “Canine morphology in the larger Felidae: implications for feeding ecology.” Biological Journal of the Linnean Society. Vol. 91, No. 4 (Aug., 2007), pp. 573-592

Christiansen, P. “Sabertooth characters in the clouded leopard (Neofelis nebulosa Griffiths 1821).” Journal of Morphology, Vol. 267, No. 10 (Jul., 2006), pp. 1186 – 1198

Christiansen, P. and Wroe, S. “Bite Forces and Evolutionary Adaptations to Feeding Ecology in Carnivores.” Ecology, Vol. 88, No. 2 (Feb., 2007), pp. 347–358

Cope, E.D. “The Amblypoda (Continued).” The American Naturalist, Vol. 19, No. 1. (Jan., 1885), pp. 40-55.

Cope, E.D. “The Tertiary Marsupialia.” The American Naturalist, Vol. 18, No. 7. (Jul., 1884), pp. 686-697.

Emerson, S.B., and Radinsky, L. “Functional Analysis of Sabertooth Cranial Morphology.” Paleobiology, Vol. 6, No. 3. (Summer, 1980), pp. 295-312.

Leutenegger, W., and Kelly, J.T. “Relationship of sexual dimorphism in canine size and body size to social, behavioral, and ecological correlates in anthropoid primates.” Primates, Vol. 18, No. 1 (Jan., 1977), pp. 117-136

Lucas, P.W., Corlett, R.T., and Luke, D.A. “Sexual dimorphism of tooth size in anthropoids.” Human Evolution Vol. 1, No. 1 (Feb., 1986), pp. 23-39

Marsh, O.C. “The Fossil Mammals of the Order Dinocerata.” The American Naturalist, Vol. 7, No. 3. (Mar., 1873), pp. 146-153

Martin, L.D., Babiarz, J.P., Naples, V.L., and Hearst, J. “Three Ways To Be a Saber-Toothed Cat.” Naturwissenschaften, Vol. 87, No. 1 (Jan. 2000), pp. 41-44

McHenry, C.R., et al. “Supermodeled sabercat, predatory behavior in Smilodon fatalis revealed by high-resolution 3D computer simulation.” PNAS, Published online before print October 2, 2007

Salesa, M.J., et al. “Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite.” Zoological Journal of the Linnean Society, Vol. 144, No. 3, (Jul., 2005) pp. 363-377

Salesa, M.J., et al. “Inferred behaviour and ecology of the primitive sabre-toothed cat Paramachairodus ogygia (Felidae, Machairodontinae) from the Late Miocene of Spain” Journal of Zoology, Vol. 268, No. 3 (Mar., 2006), pp. 243-254

Simpson, G.G. “The Function of Saber-Like Canines in Carnivorous Mammals.” American Museum Novitiates, August 4, 1941

Therrian, F. “Mandibular force profiles of extant carnivorans and implications for the feeding behaviour of extinct predators.” Journal of Zoology, Vol. 276, No. 3 (Nov., 2005), pp. 249-270

Therrian, F. “Feeding behaviour and bite force of sabretoothed predators.” Zoological Journal of the Linnean Society, Vol. 145, No. 3 (Nov., 2005), pp. 393-426

Van Valkenburgh, B., and Molnar, R.E. “Dinosaurian and mammalian predators compared.” Paleobiology, Vol. 28, No. 4 (Dec., 2002), pp. 527–543

Walker, Alan. “Mechanisms of honing in the male baboon canine.” American Journal of Physical Anthropology, Vol. 65, No. 1 (?, 1984), pp. 47 – 60

Wroe, S., McHenry, C., and Thomason, Jeffery. “Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa.” Proceedings of the Royal Society B, Vol. 272, No. 1563 (Mar., 2005), pp. 619-625

Cool Animal Meme

An interesting animal I had

Chase accompanying me on the couch

I’ve had a number of pets over the years (mostly lizards, frogs, and fish), but the most “interesting” animal I’ve ever kept is one of the cats that has been living in my apartment since this time last year. Born in 2000, Chase the cat was born to a feral mother but taken in by a large family who were friends of the woman who was later to become my wife. In 2005, when I came into the picture and visited the family now and then, I would find Chase and pet him for a little while, although it seemed Chase was a little neurotic. Eventually the family got a poodle, and the poodle decided it liked to play with Chase (I don’t think I need to tell you how Chase felt, being swatted at by a big black dog), and something had to change. So my wife and I took Chase in, but he’s definitely a strange cat . I’m the only person he is affectionate towards, so if his food bowl is empty while my wife is home he won’t let her know, but the moment I walk in the door he runs to his food bowl and starts crying. If you ever meet me, you’re likely to see little white hairs all over my clothes despite my best attempts to remove them, as well; Chase sheds nearly constantly, to the point where I wonder why he’s not bald by now. Chase also enjoys foods I didn’t think any cat liked; he’ll eat watermelon, grapes, and duct tape (although I’ve prevented this whenever he’s tried). He also licks windowsills for fun and likes to stand in front of the AC at night, but to prevent myself from going into a long post about my cat’s strange behavior Chase is definitely the most interesting animals I’ve ever had.

An interesting animal I ate

Do all the spiders I’ve probably eaten in my sleep count? My family was not especially interested in exotic dishes, so meat usually equaled chicken, turkey, or beef (sometimes fish). I did try escargot once, but the most interesting animal I’ve ever eaten will probably be a mystery to me (it was likely found inside a hot dog casing).

An interesting animal in the Museum

Just one? At the moment I would pick Amphicyon, one of the “bear dogs” of the Oligocene-Miocene (many being found in North America). I have another picture, which is unfortunately on another computer, of the skeleton caught in the light of a fading winter day, the light glinting off the teeth of the specimen pictured below. As some others have mentioned in previous comment threads, there’s little doubt that skeletal remains of this animal gives a few children nightmares.

The crushing jaws of Amphicyon.

An interesting thing I did with or to an animal

Two summers ago I went to Ocean City, Maryland with Tracey to go shark tagging. Although I was seasick for most of the trip, I did catch a juvenile Dusky Shark that I helped to tag and release. Hopefully I’ll have some more interesting animal encounters in the future that are a bit less traumatic for the creature.

An interesting animal in its natural habitat

I actually haven’t seen that many exciting animals in their “natural habitat.” Growing up in suburbia, squirrels, chickadees, white-tailed deer, and the occasional opossum or raccoon were the most I could hope for. Even now, most of the wildlife I photograph is confined to zoos, but I definitely want to see as many of the big cats as possible in the wild. Actually, one of my goals is to get to the Okovango Delta in Botswana, either to study or to merely photograph and observe, as I’m very much interested in how populations of animals there differ from populations elsewhere in Africa. Maybe someday…

I was tagged by Bora, and I tag Julia, Greg, Neil, Zach, and Kate (but feel free to pick it up if you wish to do so).

A day at the fair Faire

Here are some of the aforementioned shots from the NY Renaissance Festival in Tuxedo, NY.

When we pulled in to the lot, we saw this oddly creepy dish. The set up of the area suggested that it may have once served as a small airport, but my friends and I had no idea what the dilapidated structure was used for.

It wouldn’t be a day at the Renaissance Festival without a huge turkey leg. My friend Victor poses with a drumstick of the cooked dinosaur descendant.

“All I said was the earth goes around the sun…” My friend Tim, to the right, was locked up for taking a few too many hits off his flagon of mead and making a spectacle of himself.

The loathsome Sheriff of Nottingham, victim of a vicious pie-fight.

An evil witch takes control of a “pawn” during a rather unorthodox human chess game.

Robin Hood strikes a pose.

Observing the joust from atop a rock, sporting my spiffy new Skulls Unlimited t-shirt.

Tim being fawned over by the Singing Wenches. Never have I heard so many bawdy rhymes (well, other than some of the hip hop blasting out of souped-up Honda Civics along College Ave.)

Sunset, taken from the passenger seat of Victor’s car as we drove along the Pulaski Skyway just outside of Hoboken.

And for no reason, here are my cats Charlotte (the ‘lil black one) and Beatrice (the larger one, and she still needs a good home…).

Mine!

My trip to the Philadelphia Zoo this morning presented lots of great photo ops, especially in the morning. I’ll post more of the plethora (500+) of pictures I took tomorrow, but here’s a bit of a teaser. First, the three male Amur Tiger cubs born recently. The one in the middle really loved his tire;

The male Amur Leopard also was very curious about what I was doing on the other side of the glass, being much more active than on previous occasions when I have visited (expect a larger post on Amur Leopards and their plight in the near future);

New England Journal of Medicine Peddles Kitty Woo

In the latest edition of the New England Journal of Medicine, out today, there is a new “Perspective Article” entitled “A Day in the Life of Oscar the Cat.” It is clearly a puff piece, describing how a resident cat named Oscar at Steere House Nursing and Rehabilitation Center in Providence, Rhode Island curls up to patients that only have a few hours to live. The tone of the article is a bit more mystic however, suggesting that Oscar sniffs the air in order to figure out when someone’s “time” is, acting like a feline Grim Reaper (although to the best of my understanding Death does not purr and nuzzle those about to die, despite his well-known fondness of cats).

Still, the fact remains that this particular cat has attended 25 people who died shortly after Oscar curled up to them, and the cat generally ignores other people. This has led Yahoo! to put the story “Oscar the cat predicts patients’ deaths” on the front page, referring to the NEJM article as if it were some kind of new paper or research. The only real rationality in the Yahoo! story is found in this quote;

Nicholas Dodman, who directs an animal behavioral clinic at the Tufts University Cummings School of Veterinary Medicine and has read Dosa’s article, said the only way to know is to carefully document how Oscar divides his time between the living and dying.

If Oscar really is a furry grim reaper, it’s also possible his behavior could be driven by self-centered pleasures like a heated blanket placed on a dying person, Dodman said.

Still, I’m sure plenty of people will latch on to the story as proof of psychic connections with animals, ESP, and other woo despite the fact that an observation has been made but no actual research has been done. I’m not even sure if there have been any studies about how animals react towards the sick or dying, although there is plenty of anecdotal evidence to suggest that pets can pick up on signs that we may miss (my parents’ dog knew my mother was pregnant with me before she did, at least that’s how the story goes). Hopefully someone will have a look at this case and determine what is actually going on, but I have the feeling that even if such a study was undertaken many people would ignore it in preference of ideas about special connections with animals.

Update the 1st: Aydin brings up something I didn’t think about at first; maybe Oscar’s actions are a variation on those of Clever Hans the horse. For more, visit Snails Tales.

Likewise, Julia pointed me to a BBC article where it states that Oscar becomes quite upset if he’s removed from the room. This is likely an important clue, and (as morbid as it sounds) maybe there is some sort of chemical/pheromone/scent or something else about the dying that attracts this cat. It would explain the behavior of selecting and being affectionate with those who are about to pass away. Hopefully someone will look into this, and I’d love to see if a reason could be found for Oscar’s behavior.

New Jersey Animals, wild and otherwise

I’ve run into quite a bit of wildlife at relatively close range as of late, plenty of which I was able to catch on film. Given that these animals were not in cages or otherwise penned up, there was only so close I could get and most of these pictures are not very good (on some of them I just used the “easy fix” option on Kodak Gallery to brighten them, given that they were taken late in the day when there was almost no sunlight).

The first of my animal encounters occurred when petsitting in Monroe township for two poodles, one of which had cornered a groundhog (or woodchuck, if you prefer) behind a soccer net in the backyard. I was able to get pretty close before letting it out of there;

Later that afternoon I spotted a Turkey Vulture by the side of the road and pulled over to try and get a few shots. It flew on top of a telephone pole, too far away to get a decent shot. Given the amount of roadkill around here, however, I’ll probably have a few more opportunities this week;

Speaking of birds, today I went out for a mini-picnic and saw a mixed group of geese and ducks. Some were Canda Geese, others domestic varities, and one seemed like some sort of hybrid. They were in the shade and swam off (probably because of Panda, the dog I was watching, pictured below), so I wasn’t able to get much of a shot;

White-Tailed Deer are also ever-present in these parts, and this afternoon we got a visit from a doe and three fawns. I have seen does with as many as six fawns following, all in a group, so I wonder if white-tailed deer have their own equivalent of babysitters (they didn’t seem inclined to all stay together when I accidentally spooked them). And, if you haven’t read it already, see Friday’s post about how antlers and sexual selection figure into current deer evolution;

Meanwhile, I still check in and take care of the cats whenever I can. Thankfully they have not gone feral in my absence from the apartment, but Charlotte and Beatrice (who still needs a home) were quite interested in Spider-Man for some reason;

You Want This Cat

Just in case you all were wondering, we’re still fostering Beatrice, a lovely little tortie who needs a home. If you’re in or around central NJ (or even within a 2 hour drive or so) and are interested in adopting her, please let us know. Beatrice is two years old, she’s been spayed, she’s had all her shots, she gets along well with other cats (she used to be feral and lived with other cats), and she does not act aggressively towards people at all. If you’d like to know more, you can e-mail me at evogeek AT gmail DOT com.

Sweet Kitty Needs a Home!

For anyone nearby (in the New Jersey area) who is looking for a sweet feline friend, Friday afternoon my wife and I took in “Lady Beatrice,” a previously feral cat who was spayed and was going to be released back outside. We were going to foster her until she found a good home, but she doesn’t get along well with out cats Chase and Charlotte, so we can’t keep her here much longer.

She’s 2 years old, a black/orange/brown tortise-shell coat, EXTREMELY friendly (she loves people and is always purring, meowing, etc.), has had all her shots, is spayed, and free to a good home. I’d really love to keep her longer but my cats have been on edge ever since Friday and I can’t keep her in the bathroom indefinitely, so if you’re interested please e-mail me at evogeek AT gmail DOT com or comment on this post. I have pictures should you want to have a look at her as well.

My crazy cats

Being that I’m uploading oodles of photos, I figured I might as well include two of the cats that my wife and I share our apartment with.

Charlotte has now made a habit of attacking the broom whenever I sweep the kitchen. Normally she’s not interested in dust bunnies, but she always likes to “help” when I have a job to do.

Our cats don’t hate each other, but the older one Chase is a bit of a grump. Therefore it was a bit of a rare occassion to see them sleeping peacefully together, although a few minutes later Charlotte wore out her welcome.

A mummy dinosaur?

I know some of you must be thinking “What the hell? That’s not funny.” Well, maybe it isn’t, but I tried my best *runs off and sobs quietly* but I thought I would use an lolcat as an intro to a little paleontology lesson. Maybe this picture will help make some sense out of things;

Taken from “Integument of the iguanodont dinosaur Trachodon“, Memoirs of the AMNH ; new ser., v. 1, pt. 1-2.

If you visit the American Museum of Natural History’s fourth floor, look carefully around the mounted Anatotitan bones; near the looming, tail-dragging skeletons should be a case revealing the remains of a “mummy” dinosaur, the one pictured above. This important fossil was discovered by George Sternberg (son of Charles Sternberg, who accompanied his son on this trip and oversaw the excavation) in 1908 in Converse County, Wyoming. Osborn’s paper, however, notes an even earlier discovery of such skin impressions from a hadrosaur;

First among these integument specimens to be discovered was the famous type of Trachodon mirabilis Cope (Amer. Mus. No. 5730), found by Dr. J. L. Wortman in 1884 and now mounted in the American Museum of Natural History as part of the Cope Collection. This animal is said by Dr. Wortman to have been surrounded by a natural cast of its epidermal impressions, which unfortunately were largely destroyed or lost in the removal of the skeleton from its surroundings. There are only three patches of epidermis remaining from the tail of this specimen.

At the time both Cope’s specimen and Sternberg’s specimen were attributed to the genus Trachodon, but Trachodon is no longer accepted as valid today, but whether the fossils we’re talking about here should be called Anatotitan or Edmontosaurus still seems to be debatable. The Wikipedia entry for Anatotitan sums things up fairly well in its “Taxonomic History” section for this dinosaur, a more in-depth discussion needing a full blog post of its own. For our purposes here, however, I will be calling the dinosaur Anatotitan as that is what the AMNH (where the specimen in question is housed) has deemed fit to call it.

In any event, the “integument” of this specimen (unlike Wortman’s) was carefully preserved by the Sternbergs, allowing scientists to get a detailed look at what the skin of this particular dinosaur looked like in places. While the “tubercles” (or scale-like structures) of this fossil are certainly of interest (I suggest you take a look at the paper and its many photographs/plates for yourself), what most interested me was Osborn’s discussion of the hand of Anatotitan. In the section “Epidermal Sheathing of the Manus,” Osborn describes how the forelimbs of Anatotitan were found encased in a kind of sheath or mitt, which Osborn also mentions preserved the digits were apparently “connected by an integumentary web which was developed even more prominently than in the swimming birds.” This seemed to fit in with the contemporary hypothesis that at least some hadrosaurs were primarily aquatic, and Osborn writes;

The presence of the broad marginal web of the manus and absence of enlarged tubercles either on the dorsal or ventral surface certainly tends to support the theory of a swimming rather than of a walking, or terrestrial function of the fore paddle.

The “mitt” of Anatotitan. Taken from “Integument of the iguanodont dinosaur Trachodon“, Memoirs of the AMNH ; new ser., v. 1, pt. 1-2.

Indeed, because the forelimbs didn’t seem to seem to show a coarser or denser skin covering (and because they were held together in a “web”), it was assumed that such a limb would not work well on land and therefore it must have been an adaptation for life in the water, perfectly consistent with the image of hadrosaurs already in mind. Indeed, the “web” between the fingers was really no such thing, being that webbing exists between the toes of modern birds, not as an enclosing mitten. If the forelimbs really were paddles, we would also have to wonder why the back legs were not equipped with webbing or some other structure to enlarge their surface area during a stroke, especially being that they would be much more powerful and important for swimming. This is all easy for me to say nearly 100 years after this paper was published however, being that I have available to me much more information than Osborn did when he wrote the paper. The superficially “duck-like” appearance of hadrosaurs is hard to overcome, however, and many are still shown slogging through swamps in relatively recent works of paleo-art. For a much more in-depth discussion of hadrosaurs and what their skeletons tell us about their habitats, I heartily recommend Bob Bakker’s The Dinosaur Heresies.